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Home , Cuche Formation, Cyclopteris, Floresta Formation, La Quinta Formation

Geol. Mag. 137 (3), 2000, pp. 257–268. Printed in the United Kingdom © 2000 Cambridge University Press 257 from , with discussion of their geological and palaeogeographical context

CHRISTOPHER M. BERRY*, EDUARDO MOREL† , JAIRO MOJICA‡ & CARLOS VILLARROEL‡ *Department of Earth Sciences, University of Wales Cardiff, P.O. Box 914, Cardiff CF10 3YE, Wales, UK †Departamento de Paleobotánica, Museo de la Plata, Paseo del Bosque s/m, 1900 La Plata, Argentina ‡Universidad Nacional de Colombia, Departamento de Geociencias, Apartado 14490, Bogotá, Colombia

(Received 24 June 1999; accepted 25 January 2000)

Abstract – are described from the , Eastern Cordillera, Colombia in the area of Floresta. Those identified as Colpodexylon cf. deatsii Banks and cf. sp. suggest an earliest Late Devonian () age for the formation. These or similar taxa are also found in contemporaneous deposits in western , and other elements of the Venezuelan flora are found in a geographically intermediate locality. All three Devonian plant localities in the northwest of are within the Colombian Eastern Cordillera and its northern extension, the Venezuelan Perijá Range, an area that has been integrated as a part of the so-called ‘Eastern Andean Terrane’ or ‘Central Andean Province’, supposedly accreted to the autochthonous block of the Guyana Shield during the early or before. Although both invertebrates and plants from this terrane have strong affinities to North American and European assemblages, and might be interpreted as implying a Laurussian origin for the Eastern Andean Terrane, the evidence is not yet unequivocal, with some authors postulating an in situ development of this province.

exposures (for the most part Middle Devonian) are 1. Introduction sparse within this area, and occur generally in massifs Records of vascular plants are rare in Devonian sedi- with to ?Early Palaeozoic crystalline ments of South America. The most comprehensive cores. The Floresta Massif is composed of a crys- reports have been from the Middle and Upper talline core unconformably covered by Devonian sedi- Devonian of western Venezuela (Berry, Edwards ments that include the El Tibet, Floresta and Cuche and co-authors) and the /Lower Devonian formations. The El Tibet Formation has a restricted of Bolivia/Argentina (Morel, Edwards & Iñigez distribution and thickness ranging up to 425 m (M. G. Rodriguez, 1995). This scarcity of evidence has Paz & A. D. Urrutia, unpub. Trabajo de Grado, frustrated attempts at global phytogeography during Departamento de Geociencias, Universidad Nacional these times (Edwards & Berry, 1991). de Colombia, Bogotá, 1996). It is composed mainly of The present report concerns fossil plants from fine- to coarse-grained quartzitic and kaolinitic sand- Devonian strata of the Cuche Formation, outcropping stones, deposited essentially in a continental environ- in the Eastern Cordillera, Colombia around Floresta ment (braided streams) and coastal to subcoastal in the Department of Boyacá (Figs 1, 2), and is com- conditions. On the basis of a spore assemblage, plimentary to work on fossil vertebrates in the same Grösser & Prössel (1994) defined an age for region (Janvier & Villarroel, 1998). this unit. The field observations and collection of the samples The overlying (Olsson & Caster, were accomplished by JM and CV, further samples 1939) contains fossil invertebrates (, trilo- collected by Professor Medardo Chacón, and study of bites, bryozoans, gastropods, etc.) as described by the fossil plants was undertaken by CMB and EM. Caster (1939), Royo y Gómez (1942), Morales (1965), and fish remains of rhenanid and arthrodire type (Janvier & Villarroel, 1998). The fossil content indi- 2. Geological background and localities cates a Middle Devonian age for the Floresta As shown in the publications of Barrett (1988) and Formation. Forero (1991), known outcrops of sedimentary The Cuche Formation (Botero, 1950) is the Devonian rocks in Colombia are completely restricted youngest of the Palaeozoic sedimentary sequences to a region comprising the Eastern Cordillera, its developed in the Floresta Massif. It represents a northward extension in the Sierra de Perijá and the regressive unit deposited in a transitional environment upper Magdalena Valley. However, the Devonian that includes tidal to supratidal (Lower Member) and lagoonal siltstones containing a fossil * Author for correspondence: [email protected] assemblage of vertebrates, invertebrates and plants

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Figure 2. Location map of the two fossil localities men- tioned in this report: (1) Puerto Arepas; (4) Quebrada Potrero Rincón. Modified after Mojica & Villarroel, 1984.

Figure 1. Devonian outcrops yielding fossil plants in Colombia and northwestern Venezuela (stars) in relation to known in the adjoining Santander Massif, studied in selected terranes. AUT= Autochthonous Terrane (Guyana detail by Rabe (1977). Shield), EAT = Eastern Andean Terrane (including Sierra de Mojica & Villarroel (1984) provided a list of plant Perijá and Eastern Cordillera). Modified from Restrepo & Toussaint, 1988. fossils from the Cuche Formation, including the gen- era Psilophyton Dawson emend. Banks, Leclercq & Hueber and Ginkgophyton sensu Beck. Other fossils include the Welleria Ulrick & Bassler and which reflect the transition from typical shallow fish remains. On this basis Mojica & Villarroel (1984) marine conditions to a nearshore terrestrial domain. assigned an age from Middle Devonian to Early This unit is made up of beds of sandstones and silt- to the Cuche Formation. However, stones, which have a concordant and transitional rela- recent study of the vertebrate fauna has allowed tionship with the underlying Floresta Formation. It restriction of the time of deposition to the Late consists of a Lower Member (140 m) of sandstones Devonian Epoch, specifically to the Frasnian (Janvier and an Upper Member (250 m) in which there are & Villarroel, 1998). thick beds of red, violet and ochre sandstones and silt- The plant fossils studied in this paper include the stones. The Lower Member contains many small, same specimens reported by Mojica & Villarroel unidentifiable fragmentary plant remains, but the (1984) as well as others, collected later by JM and CV. Upper Member contains several beds with well-pre- The fossils described here were collected near served fossil plants. The has Floresta, in the Eastern Cordillera (Fig. 1). Most been interpreted as shallow water, of varying energy, derive from the section through the upper part of the with occasional subaerial exposure (Mojica & Floresta Formation and the Cuche Formation in the Villarroel, 1984). The Cuche Formation is overlain dis- upper part of Quebrada Potrero Rincón (Fig. 2, local- cordantly by a thick sequence of (conglomer- ity 4) described by Mojica & Villarroel (1984, their figs ates, sandstones and siltstones) resembling the Upper 3, 4). A complimentary stratigraphic section was pro- Jurassic Girón Formation (Hettner, 1892). After the vided by Janvier & Villarroel (1998, their fig. 2). The deposition of the Cuche Formation and before the best preserved and most informative plant specimens accumulation of the Girón-like red beds, the Floresta are from assemblage III (Mojica & Villarroel, 1984, Massif was uplifted leading to no deposition or ero- their fig. 3), derived from a locality made up of sandy sion of the Carboniferous and sequences of a yellow/brown colour which also contain

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fish scales and bones (Janvier & Villarroel, 1998, separated by a distance of approximately 1.6 mm. Leaf locality PR-1). Plant fossils include the specimen of bases alternate in successive gyres. Leaf bases are also Colpodexylon, the unnamed trimerophyte derivative, visible on the left-hand margin of the stem where they cf. Archaeopteris and some of the leafy fossils. The are thickened compared to the proximal leaf lamina, horizon is in the middle of the Upper Member of the appearing decurrent in compression beside the stem. At Cuche Formation. the widest part of the remaining stem there are a maxi- A single large leaf, Mch-019-e, was recovered from mum of six leaf bases visible in any gyre, implying a min- the same section in Quebrada Potrero Rincón, imum of 12 in any complete rotation. towards the bottom of the Lower Member, where rain drop impressions (‘gotas de lluvia’) are recorded just Leaf morphology. A single basal portion of a leaf below thick beds of reddish sandstones (Mojica & going down into the matrix below the stem has a width Villarroel, 1984, fig. 3). of 1.0 mm. Other examples uncovered compressed Other large leaves have been found in the site named beside the stem are much thinner (0.4–0.6 mm), this ‘Puerto Arepas’ on the south side of the road from being due to compaction and folding of the lamina. to , just below an angular un- Four leaves have been uncovered and are com- conformity with the conglomerates of the Early paratively well preserved. Their measurements are Formation. This locality is recorded in Table 1. Lengths of leaves and segments marked as 1 on Figure 2. These fossils are preserved in are minimum only, as the tips are so delicate it is diffi- red micaceous . cult to uncover them, and most are incomplete. The leaves have a maximum observed length of 21.9 mm. Uncovering the four well-preserved leaves revealed 3. Methods them to be divided. Careful removal of one segment of Plant megafossils were uncovered using needles the fourth leaf (segment 1, Figs 3b–d, 4a,b) revealed (dégagement: Fairon-Demaret, Hilton & Berry, 1999). two further segments below (segments 2, 3). The leaves They were photographed using a Canon F-1 camera, are therefore trifurcate. A reconstruction is given in macro lens and extension tubes where necessary. All Figure 4d. Two short segments are arranged either side photographs were taken dry using crossed polarizing of a medial longer segment. They have acute tips filters to enhance contrast. Palynological preparations where preserved. using standard techniques showed the matrix to be barren of palynomorphs. Identity and comparisons. Of the well known species of The fossils are retained in the collections of the Devonian herbaceous lycopsids, only Colpodexylon Departamento de Geociencias, Bogotá. Banks (1944) has leaves which are divided into three segments in such a manner. Literature on this genus was recently reviewed by Berry & Edwards (1995). Leaves of 4. Palaeobotanical descriptions Minarodendron (Li, 1990) have three segments of which Colpodexylon cf. deatsii Banks the central segment is perpendicular to the other two Figures 3a-d, 4 and differs also in that the margins are serrate. Colpodexylon is well known from the Devonian Material. Specimen no. CCHE-IX-86-023; locality 4, strata of New York State (Banks, 1944; Grierson & assemblage III. Banks, 1963). Two species (C. deatsii, C. trifurcatum) were originally distinguished by Banks (1944) on the Stem morphology. The stem is 36 mm long and 9.3 mm basis of measurements of leaf and sporangial dimen- in exposed width (Fig. 3a), although it appears not to be sions. Edwards & Benedetto (1985) described a new completely preserved on the right-hand side. It is pre- species, C. cachiriense, with shorter leaves and bases served as an impression, with a small amount of organic which were not contiguous, from western Venezuela. material remaining. The stem surface is dark brown in A species with proportionally shorter segments distal colour. Leaf attachments are represented by inverted v- to the trifurcation was described by Schweitzer & shaped depressions arranged in more or less flat rows Cai (1987) from the of southwestern

Table 1. Measurements of dimensions (in millimetres) of leaves of Colpodexylon cf. deatsii from Colombia

Total length Length to Width below Length median Width median Length lateral Width lateral Leaf of leaf trifurcation trifurcation segment segment segment(s) segment(s)

a 21.9 12.5 0.46 9.4 0.46 2.9 0.17 b 0.41 0.31 1.65 0.15 2.9 0.12 c 0.39 0.34 2.9 0.15 d 0.56 0.31 0.17

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Figure 3. Plant fossils from Quebrada Potrero Rincón. (a–d) Colpodexylon cf. deatsii Banks, CCHE-IX-86-023. (a) View of fos- sil after extensive dégagement; (b–d) three views of second leaf from top during dégagement showing three distinct segments (1, 2, 3). (e–i) Unnamed trimerophyte derivative. (e) Close-up of paired sporangia showing beak and dehiscence line, constriction at base of sporangium (arrowhead), specimen no. ICN-PI-0268-1; (f) fertile terminations of axes with more rounded sporangia probably arranged in pairs (e.g. arrow), unnumbered specimen; (g) fertile termination of axes, specimen no. ICN-PI-0268-1; (h) fertile branching axes, specimen no. ICN-PI-0380-1; (i) fertile branching axes showing bifurcate and trifurcate (arrows) branch- ing, specimen no. ICN-PI-0327. Scale bars: (a) 10 mm, (b–d) 2 mm, (e, f) 5 mm, (g–i) 10 mm.

China. More recently, new species with both short leaves on the stems and the general leaf morphology it (C. coloradense) and long (C. camptophyllum) distal is superficially most similar to C. deatsii from New segments have been described from Venezuela (Berry York State, although also close to C. trifurcatum. & Edwards, 1995). Leaves do not have the same flattened appearance as With only a single fragmentary specimen it is impos- C. camptophyllum. sible to place it in any one of these species with cer- tainty, and certainly no new species can be erected. Unidentified trimerophyte derivative However, on the basis of the close arrangement of Figures 3e-i, 5

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Figure 5. Line drawings of unnamed trimerophyte deriva- tive. (a) ICN-PI-0380-1, (b) ICN-PI-0327 (see Fig. 3h, i). Scale bar = 5 mm.

Specimen no. ICN-PI-0380-1 Figures 3i, 5b

Basally, there may be a trichotomy of the stem (Fig. 3i, lower arrow), yielding an uncovered branch (central) and a dichotomous branch bearing terminal sporan- gia (right). The main axis is 1 mm in diameter and tri- chotomizes more or less equally to give three 0.6 mm Figure 4. Colpodexylon cf. deatsii Banks, CCHE-IX-86-023. diameter axes 11 mm from the base (Fig. 3i, upper (a, b) Sequential stages of dégagement of second leaf from arrow). The left-hand axis is not preserved and the top in Figure 3a (see Fig. 3b–d); (c) outline of top leaf right-hand was only partially uncovered. The central in Figure 3a; (d) reconstruction of whole leaf. Scale bar = 5 mm. axis branches again dichotomously 6 mm from the previous branching point, and each of the daughter axes dichotomizes further. Beyond this, pairs of termi- Material. Four slabs bear fragments of an as yet nal sporangia are visible, although their preservation is unidentified trimerophyte derivative for which, indistinct, following one or a number of more closely because of a lack of counterparts and of preserved spaced dichotomies. anatomy, it is impossible to make a satisfactory description or make taxonomic deductions. However Specimen no. ICN-PI-0327 they probably all belong to the same species. They Figures 3h, 5a were previously assigned to Psilophyton and probably P. microspinosum by Petriella (in Mojica & Villarroel, This specimen has a maximum length of 41 mm and is 1984). Dégagement revealed far more of the branch- 0.6 mm in diameter at the base. The distantly spaced ing pattern than had been observed on the surface by basal branching points suggest a monopodial arrange- previous observers. All these fossils derived from local- ment. Branching is not well-enough preserved to ity 4, assemblage III. determine if it is dichotomous or trichotomous.

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A number of close dichotomies distally yield paired traits, namely centrach terrete primary xylem strand, terminal sporangia. pitted tracheids and dichotomous division of the xylem strand into distal branches, are not presently Specimen no. ICN-PI-0268-1 known from Middle Devonian plants. The only Figure 3e, g Middle Devonian species named Psilophyton, P. kräuseli, was established by Obrhel (1959, 1961) for a Dichotomous axes bear distally terminal sporangia. small number of Middle Devonian specimens from These are of 2.0–2.8 mm in length and 0.9–1.2 mm in Bohemia. The plant is spiny, yet lacks preserved width and apparently flattened. The arrangement of anatomy, and the single fertile specimen is not well the sporangia is not clear with the material to hand. enough preserved to demonstrate the exact arrange- They lie in different orientations and a lack of coun- ment of sporangia. Although dichotomous branching terpart prevents meaningful dégagement. The most prevails in the genus Psilophyton, trifurcate branching obvious disposition is that they are arranged in pairs is known from Lower Devonian P. dawsonii (Banks, (Fig. 3g, furthest right). Others may contradict this Leclercq & Hueber, 1975), where sometimes an assertion; for example, those in Figure 3g, top left aborted third trace is present at a branching point. (enlarged in Fig. 3e) strongly suggest dehiscence has However the alternating nodal angle of the main axis occurred and the valve has broken into two more or in our largest specimen, giving a zigzag effect, hints at less equal halves. It is possible that this second a higher level of anatomical complexity. For example, arrangement may take on the appearance of the first if ribbed primary xylem columns are responsible for an compressed at a different angle. It is also possible that identical zigzag, trifurcate branching pattern in those illustrated in Figure 3e might represent two anatomically preserved Stenokoleos holmesii (Matten, sporangia arranged as a pair, with straight margins 1992). However the morphology of plants in the adjacent and curved outer margins. One sporangium Stenokoleales remains unknown. Unfortunately our shows a possible constriction in the axis at its base, specimens are not large enough to determine if there is which may demonstrate a specialized abscission layer a regular and repetitive orientation of the lateral (Fig. 3e, arrowhead). branches as one might expect for a ribbed vascular sys- tem of the main stem, or a random orientation that Unnumbered specimen one might expect from a Psilophyton-like anatomical Figure 3f organization. Unpublished plants with a similar branching pat- Two dichotomous trusses lying close together on the tern are known from Middle or lowermost Upper slab have several sporangia visible, arranged in pairs Devonian assemblages from Venezuela, yet have more with straight margins on the inner face of the pair. The complex sporangial clusters reminiscent of aneuro- outer margins of the sporangia of this specimen are phytalean progymnosperms. more strongly convex than in the other examples. The sporangia are 1–1.3 mm in length and 0.5–0.8 mm cf. Archaeopteris Dawson wide. Figure 6a, b

Discussion and comparisons. Small dichotomizing Material. These fossils derive from locality 4, assem- branched axial plant fossils reminiscent of the Early blage III. Devonian trimerophyte genus Psilophyton are com- monly found in Middle and lowermost Upper Specimen no. CCHE-IX-86-026 (a) Devonian sediments. They are often attributed to the Figure 6a form genus Psilophytites (Høeg, 1952) when sterile and bearing small spines. Numerous leaves are present on this slab. The most Although the exact disposition of the sporangia informative are the four inserted on the right-hand (paired or not paired?) remains unresolved, the margin of an axis 3 mm wide, spaced 20 mm apart. arrangement in pairs of sporangia in the unnumbered The surface of the axis is marked by small transversely specimen (above) is preferred to a single dehisced spo- oriented ornament. The leaves have narrow bases, are rangium because the single sporangium would have about 33 mm in length and 20 mm maximum width, to be globose in order to achieve the morphology and have an entire margin. They are of an overall fla- pictured after a median dehiscence (Fig. 3f). belliform morphology. Venation is dichotomous, with The fossils described above are likely to derive from the distal veins curved away from the midline of the trimerophyte ancestors because of the probably paired leaf towards the margins. Other fragments on the spec- terminal sporangia and general morphology typical of imen suggest leaves which were considerably larger. this . No Middle or Upper Devonian plant has Elongate 1.5 mm long sporangia are also present, but been shown to have both morphological and anatomi- not in attachment to other organs. Larger stems with a cal characteristics of the genus Psilophyton. The latter similar ornament are visible.

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Figure 6. Plant fossils from Quebrada Potrero Rincón. (a, b) cf. Archaeopteris Dawson. (a) Three leaves attached to axis and a fragmentary fourth (lowest) also believed attached, CCHE-IX-86-026; (b) five leaves attached to axis, ICN-PI-0378-1; (c) leaf fossil, ICN-PI-0292-2. Scale bars = 10 mm.

Specimen no. ICN-PI-0378-1 parts are in evidence which would allow unambiguous Figure 6b assignment to this genus. Delimitation of Archaeopteris species based on frag- This axis, of 2 mm width, bears five leaves of 22 mm mentary specimens is neither well founded nor wise maximum length in an apparent alternate arrange- (Gensel & Barnett-Lawrence, 1996). There is consider- ment. They are inserted at approximately 45° on nar- able polymorphism and variation in leaf size within row bases, but enlarge to a maximum width of 20 mm individual fronds (Kenrick & Fairon-Demaret, 1991) distally. Proximal margins are more or less straight and presumably within branches and on individual and the leaves probably entire. Venation is dichoto- trees. mously branched. Our Archaeopteris is most similar to A. halliana from North America and A. roemeriana from Belgium Comparison and identity. These specimens from the (the former synonymized with the latter by Kräusel & upper fossiliferous horizon, with a number of entire Weyland (1941)) in terms of leaf morphology, vena- leaves inserted upon axes with a transverse ornament tion and overlap of the leaves. However leaves of A. can only, in the context of Devonian sediments, be halliana were defined as less than 20 mm by Kräusel & attributed to the genus Archaeopteris Dawson. They Weyland (although leaves ‘slightly over 2 cm long’ are therefore believed to be the leafy branches of a were included in the species by Gensel & Barnett- large arborescent progymnosperm. No attached fertile Lawrence (1996), based on material from the Frasnian

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of Gaspé, Canada). The attached leaves are up to Such leaves are often referred to Psygmophyllum or 33 mm in the Colombian specimens, and so may be Flabellifolium (see e.g. Gensel & Barnett-Lawrence, more comparable to A. obtusa Lesquereux. However, 1996). The specimen from locality 4 is better preserved with so few fragmentary, sterile specimens, any but is of squatter outline. We do not feel compelled to attempt to assign these fossils to a species is futile. assign generic or specific names to this material; rather Archaeopteris is widespread in the sediments of Late its occurrence suggests the presence of Archaeopteris- Devonian age, known also from high and low latitudes type plants in the Lower as well as Upper Member of of (South Africa, Australia) as well as the Cuche Formation. New York State, Ellesmere Island, western Europe (Laurentia), Siberia and China. 5. Stratigraphic implications Leafy fossils In New York State, Colpodexylon is restricted to the Givetian and lowermost Frasnian deposits. It is also Specimen no. MCH-019-e known from Venezuela, Vietnam, and south and Material. Rain drop horizon, lower level of Lower northwestern China during the same time (Berry, Member of Cuche Formation, Quebrada Potrero 1997). Archaeopteris species with entire leaves are first Rincón. found in lower Frasnian deposits of Gaspé, Canada (Gensel & Barnett-Lawrence, 1996; McGregor, 1996) Description. This fossil is an incomplete leaf, faintly and continue at least to the latest part of the Devonian preserved, of maximum length 63 mm. At the most Period (Fairon-Demaret, 1986). Trimerophyte deriva- proximal preserved point it measures only 4 mm wide, tives superficially similar to ours are known from 20 mm more distal it is 12 mm wide, but it widens Lower, Middle and lower Upper Devonian sediments quickly to a maximum preserved width of 43 mm, it and have less stratigraphic significance. being flabelliform in outline. The leaf veins are indis- Devonian plant biostratigraphy is not yet an exact tinctly preserved and curve outwards towards the dis- science (Edwards & Berry, 1991; Edwards, Fairon- tal margins. A second specimen (unnumbered) in a Demaret & Berry, in press). However the co- slightly coarser red matrix suggests that the leaves were occurrence of Colpodexylon and Archaeopteris in inserted acutely and possibly spirally upon an axis, but assemblage III suggests a Late Devonian (earliest the axis itself is missing and the three leaves are more Frasnian) age for the middle of the Upper Member of fragmentary than that described above. Both fossils the Cuche Formation. The presence of large leaves are poorly and indistinctly preserved as impressions near the base of the Lower Member suggests it is also lacking organic carbon. of the same age. Archaeopteris and Colpodexylon co- occur in Livingstone Quarry, Oneonta Formation Specimen no. ICN-PI-0292-2 (lower Frasnian) in New York State (Banks, Grierson Figure 6c & Bonamo, 1985).

Material. Locality 4, assemblage III. 6. Comparison with other contemporaneous South Description. This flabelliform leaf has a minimum American assemblages proximal width of 6 mm and a maximum width of 6.a. Venezuela 52 mm some 20 mm from the base. Its maximum length is 45 mm. Veins dichotomise distally (basally The most diverse, best known, Devonian floras of they measure 0.15 mm to an occasional 0.3 mm) and Middle and Late Devonian age in South America are widen distally where they curve away from the midline those from the Campo Chico Formation in the Río of the leaf towards the distal lateral margins. The leaf Cachirí area of the Sierra de Perijá, northwestern is entire, as far as can be judged, with concave margins Venezuela. The Lower Member of the Campo Chico proximally and convex distally, and its width to length Formation contains a diverse flora composed of zos- ratio is considerably higher than those discussed in the terophylls (cf. Sawdonia; Serrulacaulis cf. furcatus, preceding paragraph. Berry & Edwards, 1994), herbaceous lycopsids (Leclercqia cf. complexa, Berry, 1994; Haskinsia sagit- Identity and comparisons. The isolated leafy fossils tata Edwards & Benedetto, 1985; H. hastata Berry & from the lower red beds are very large, comparable Edwards, 1996a; Colpodexylon cachiriense Edwards & with the largest pinnules of the sterile type specimens Benedetto, 1985; C. coloradoense, C. camptophyllum of Archaeopteris obtusa (Lesquereux, 1880). Their Berry & Edwards, 1995; Gilboaphyton griersonii Berry morphology and insertion is comparable to other & Edwards, 1997), iridopteridaleans (Anapaulia illustrated examples of A. obtusa from the Upper moodyi Berry & Edwards, 1996b; new genus), cladoxy- Devonian sediments of Gaspé (Cyclopteris obtusa of lopsids (Wattieza sp.), progymnosperms (Rellimia sp.), Dawson, 1882; Gensel & Barnett-Lawrence, 1996). a possible rhyniopsid (with S-type vascular tissues),

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and various other as yet unclassified plants. These are Dolianiti, 1957). Protolepidodendron kegeli has pre- believed to be of Givetian (late Middle Devonian) age served leaves which were reported to be similar to (Berry, Casas & Moody, 1993) but may be also of tuning forks. ‘Archaeosigillaria’ picosensis was a stem latest or early Frasnian age. impression with characteristic stem-surface features. Recently, in the Upper Member of the Campo Lastly, Kraüsel & Dolianiti reported stems super- Chico Formation, a more restricted flora has been dis- ficially similar to lycopsids with randomly arranged covered, composed predominantly of two species of ‘leaf scars’ under the name Palaeostigma sewardi. Archaeopteris, one with large, entire, flabelliform These last two taxa are also known from South Africa leaves, and a second with smaller, more dissected (Anderson & Anderson, 1985; Chaloner et al. 1980). leaves superficially similar to those of A. notosaria from South Africa (Anderson, Hiller & Gess, 1995). 7. Geological context The two Colombian localities (Floresta, Pamplona) 6.b. Colombia demonstrate the continuation of the flora known to A small florule of Middle Devonian age was reported have flourished in northwestern Venezuela during late from the Middle Devonian of Chitagá Canyon, near Middle and earliest Late Devonian times some Pamplona, northeastern Colombia (Boinet et al. 600–700 km further southeast in present-day terms 1986). Three taxa were recognized. A single specimen (Fig. 1). The Venezuelan flora, exceptional in its diver- of the herbaceous lycopsid Protolepidodendron sp. sity, most clearly resembles that of New York State requires demonstration of leaf morphology before it and western Europe. Some species are in common can be correctly identified, yet the sections through the between New York and Venezuela (Gilboaphyton leaves illustrated suggest it could belong to Haskinsia griersonii, Haskinsia sagittata/collophylla, Leclercqia or Leclercqia. A sterile axis of cf. Stockmansella langii complexa, Serrulacaulis furcatus amongst those groups has a prominent vascular strand and is superficially so far investigated in detail). There are also some identical to the ‘possible rhyniopsids’ reported (above) similarities to Belgium (e.g. Wattieza sp., Serrulacaulis, from Venezuela. A leaf of 35 mm length attributed to Leclercqia). It is not possible yet to demonstrate such Psygmophyllum cf. williamsonii is typical of disarticu- relationships with the less well preserved floras of lated Archaeopteris. Some dichotomizing axes are also Argentina and Brazil (the latter appearing to show at present. least slight affinity to South Africa). Although a dis- parity in vegetation is possible, this observation may be at least partly due to the paucity of the record in 6.c. Elsewhere in South America mid and southern South America. Away from the northwestern corner of South The similarity of Devonian invertebrate faunas of America, Middle and Upper Devonian plant fossils the Sierra de Perijá and Colombia to those of the are less well preserved, less diverse, and even less confi- Eastern Americas Invertebrate Realm is more strongly dently dated. defined (e.g. brachiopods: Benedetto, 1984; : In Argentina, lycopsids are common in sediments Scrutton, 1973; Meyerhoff et al. 1996) and, because of believed to be of Middle Devonian age, and are attrib- higher numbers of localities and better geographical uted to the genera Haplostigma and Malanzania coverage, provides a better basis currently for palaeo- (Gutierrez, 1996). These do not have well preserved geographical deliberation than plant megafossils. complex leaf morphology which precludes their being There are quite different opinions about the origin placed in genera such as Haskinsia, Colpodexylon or and tectonic relationships of the north Andean Leclercqia. Frenguellia, a generic concept developed provinces and the western pericratonic region of the from fossils from widespread localities with little age Guyana Shield during Palaeozoic times. Both regions constraint, demonstrates complex leaf morphology are now separated by a complex thrust fault system and adaxial sporangia on unmodified leaves, charac- known as the Borde Llanero Fault System, Borde ters known only from Lower, Middle and lowest Llanero Suture or Guaicaramo–Yopal Fault System. Upper Devonian lycopsids; however these plants are Some previous authors have assumed that the reported to be from the Lower Carboniferous Devonian deposits of northwestern South America (Arrondo, Cesari & Gutierrez, 1991; Morel et al. have been deposited on the margins of the Gondwana 1993). Specimens attributed to the Middle Devonian supercontinent (Barrett, 1988; Benedetto, 1984; Berry, cladoxylopsid Hyenia (Morel et al. 1993) are more 1994) and are more or less in situ. Maze (1984) sug- likely to represent erect aerial axes rather than trailing gested, based on palaeomagnetic data, that area to the rhizomes and therefore the sterile specimens cannot be west of Lake Maracaibo might be allochthonous. identified, perhaps being of progymnosperm origin Restrepo & Toussaint (1988) considered that the (Fairon-Demaret & Berry, in press). Palaeozoic to Early Mesozoic history of the A few specimens have been attributed to lycopsids autochthonous block (pericratonic region: AUT in from the Devonian of Piauí State, Brazil (Kraüsel & Fig. 1) is fundamentally different from that of the

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Eastern Andean Terrane immediately to the west whole region was part of one tectonic province or ter- (EAT in Fig. 1), and only events are rane. Moreover, the Lower Palaeozoic sediments of the common to both blocks. Consequently, Restrepo & Eastern Andean Terrane show no evidence of low Toussaint (1988) concluded that suturing occurred grade metamorphism (Jiménez & Caicedo, 1998) or of sometime during or before Jurassic times. Each of the having been highly deformed (Mojica, Villarroel & three plant-bearing Devonian deposits discussed above, Macia, 1987) by any Palaeozoic orogenesis as supposed Sierra de Perijá (Venezuela), Pamplona and Floresta by those authors favouring an allochthonous origin. (Colombia), is within the Eastern Andean Terrane. The results obtained in this work indicate that the A quite similar scenario for tectonic development Late Devonian plants described here have closer rela- during Devonian times was presented by Forero tionships to those of the Laurussian continent than (1990), who considered that the Borde Llanero Fault can yet be demonstrated for the Gondwana continent. System represents the limit between two Early However, the presence of typical Gondwanan ele- Palaeozoic provinces, the Guyana Shield (Gondwana) ments (e.g. Antarctilamna) in newly discovered verte- to the east and an allochthonous terrane (formerly a brate assemblages associated with the plants from piece of the North American continent) to the west, Colombia and Venezuela (Janvier & Villarroel, 1998; the last comprising the Eastern Cordillera, the G. C. Young, pers. comm.) suggest also a connection Magdalena Valley and the eastern part of the Central with the east part of the north Gondwana margin, Cordillera. Forero (1990) postulated that both ter- probably in close proximity to or in contact with the ranes were accreted during Late Silurian to Early Laurussian supercontinent. The origin of this dis- Devonian times (Caledonian Orogenesis) as a conse- agreement is not yet clear. Possible future discoveries quence of the collision between North and South of additional Devonian localities across Gondwana America (Gondwana). Consequently, the Devonian may help in solving this problem. sediments of both regions should lie on a Precambrian to Early Palaeozoic , and the Devonian ben- thic-faunal records on the neighbouring sides of 8. Conclusions North and South America have to be very similar. In (1) As indicated by the palaeontological results this context, Forero (1986) pointed out that during obtained in this work and that of Janvier & Villarroel Late Devonian times (Frasnian–Famennian) faunal (1998), the sedimentary rocks representing the and lithofacial similarities suggest a close geographic Devonian System in Colombia include not only the relationship between the areas encompassed by the often mentioned shallow marine sequence of the Devonian Old World Realm (North America, Europe Floresta Formation, but also fossil-bearing red beds of and northern Africa). the Cuche Formation. The plant and vertebrate con- An accretionary model resulting from progressive tent indicates a Late Devonian (Frasnian) age for the westward migration of subduction zones has been also whole Cuche Formation. proposed by McCourt, Aspden & Brook (1984), who (2) The sedimentary record of the Upper Devonian considered that during the ?Devonian a subduction in northwestern South America discussed here zone existed oceanward of the Precambrian Guyana comprises a narrow northeast to southwest belt Shield, and that the Bolo Azul and the Rosario com- (600–700 km long), restricted to the north-central plexes of the Central Cordillera are remnants of the Eastern Cordillera of Colombia and the eastern flank volcanic arc. The quartz-rich, low metamorphic grade of the Perijá Range. The above mentioned Upper Cajamarca Group (Nelson, 1957) might represent Devonian belt has not been taken into account in rela- accumulation at the same time to the east of the arc. In tively recent palaeogeographical reconstructions, e.g. this context, the Devonian sediments in the Eastern Barrett (1988). Cordillera should have been deposited more or less in (3) The flora and fauna of the Cuche Formation and situ. However, the expected Devonian dating of the other Middle and Upper Devonian sediments from Cajamarca Group and of the Bolo Azul and Rosario Colombia and Venezuela show closest similarity to complexes remains speculative and their possible time those of eastern North America and western non- relationships to the Devonian sediments of the alpine Europe. Eastern Andean Terrane have to be proven. Mojica & Villarroel (1990) have postulated an in situ Acknowledgements. JM and CV are very grateful to those development of this region, taking into account that persons of the Universidad Nacional de La Plata who first the Lower Palaeozoic of the western border of the established the importance of studying the plants of the Guyana Shield, the Llanos area, the Eastern Cuche Formation (especially Professors Bruno Petriella, Cordillera and the Magdalena Valley show a continu- Sergio Archangelsky and Oscar Arrondo) and encouraged us to conduct more detailed fieldwork and collect more spec- ous, coherent lateral facies development that reflects imens. CMB would like to thank the Geological Society, an eastern located source area and sedimentation in a London (W.G. Fearnsides Fund) for supporting fieldwork in broad, shallow platform deepening westwards towards Colombia, and Professors M. Fairon-Demaret and P.Gensel the present Central Cordillera, thus indicating that the for their helpful reviews of the manuscript.

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